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Take home messageMucilage secreted by roots and EPS produced by microorganisms alter the physical properties of the soil solution and impact the water dynamics in the rhizosphere. The high viscosity of mucilage and EPS is responsible for the formation of thin filaments and interconnected thin lamellae that span throughout the soil matrix maintaining the continuity of the liquid phase across the pore space even during severe drying. The impact of these mechanisms on plant and microorganisms needs to be explored. [ABSTRACT FROM AUTHOR]

• Soil water evaporation can be delayed by the complex biofilm producer B. subtilis. • An accelerated retreat of the evaporation front initiates decelerated water loss. • Reduced soil hydraulic conductivity and surface tension explain the process. • Modifications are result of a continuous adaptation. When soils dry, water flow and nutrient diffusion cease as the hydraulic microenvironments vital for soil life become fragmented. To delay soil drying locally and related adverse effects, bacteria and plants modify their surroundings by releasing extracellular polymeric substances (EPS). As a result, the physical properties of hotspots like biological soil crusts or the rhizosphere differ from those of the surrounding bulk soil. Specifically, the presence of EPS delays evaporative soil drying. Despite the evidence of reduced evaporation from EPS-amended soils, the mechanisms controlling soil water content dynamics remain elusive. Thus, our study aimed to elucidate the potential of bacteria to modify their local environment when exposed to oscillations in soil water content induced by evaporative drying. We incubated sand microcosms with two contrasting strains of Bacillus subtilis for one week in a flow cabinet. At the end of the incubation period, local water loss was quantified and spatially resolved using time-series neutron radiography. Strain NCIB 3610, a complex biofilm producer steadily modified soil evaporation dynamics during the incubation period resulting in a substantial delay in soil drying due to hydraulic decoupling of the evaporation front from the soil surface. Evaporation dynamics remained largely unaltered in the microcosms inoculated with the domesticated strain 168 trp+ compared to the control treatment. The mechanism of hydraulic decoupling induced by NCIB 3610 was verified by estimates of diffusive fluxes and the position of the evaporation plane in the microcosm. Additionally, the role of polymeric substances in hydraulic decoupling was confirmed by an evaporation experiment using xanthan as an EPS analogue. [ABSTRACT FROM AUTHOR]

Plant roots exude a significant fraction of the carbon assimilated via photosynthesis into the soil. The mucilaginous fraction of root exudates affects the hydraulic properties of the soil near the roots, the so called rhizosphere, in a remarkable and dynamic way. After drying, mucilage becomes hydrophobic and limits the rewetting of the rhizosphere. Here, we aim to find a quantitative relation between rhizosphere rewetting, particle size, soil matric potential and mucilage concentration. We used a pore-network model in which mucilage was randomly distributed in a cubic lattice. The general idea was that the mucilage concentration per solid soil surface increases the contact angle between the liquid and solid phases consequently limiting the rewetting of pores covered with dry mucilage. We used the Young–Laplace equation to calculate the mucilage concentration at which pores are not wettable for varying particle sizes and matric potentials. Then, we simulated the percolation of water across a cubic lattice. Our simulations predicted that above a critical mucilage concentration water could not flow through the porous medium. The critical mucilage concentration decreased with increasing particle size and decreasing matric potential. The model was compared with experiments of capillary rise in soils of different particle size and mucilage concentration. The experiments confirmed the percolation behaviour of the rhizosphere rewetting. Mucilage turned hydrophobic at concentrations above 0.1 mg/cm 2 . The critical mucilage concentration at matric potential of −2.5 hPa was ca. 1% [g/g] for fine sand and 0.1 % [g/g] for coarse sand. Our conceptual model is a first step towards a better understanding of the water dynamics in the rhizosphere during rewetting and it can be used to predict in what soil textures rhizosphere water repellency becomes a critical issue for root water uptake. [ABSTRACT FROM AUTHOR]

Plant roots and bacteria are capable of altering their local soil environment by releasing a diverse, highly polymeric blend of substances (e.g. extracellular polymeric substances EPS and mucilage). Despite the importance of EPS and mucilage for plants and microorganisms is well accepted, the physical mechanisms by which EPS and mucilage interact with the soil matrix and determine the soil water dynamics remain unclear. High-resolution X-ray CT revealed that upon drying in porous media mucilage and EPS form one-dimensional filaments and two-dimensional interconnected structures spanning across multiple pores. Unlike water, these mucilage and EPS structures connecting soil particles did not break up upon drying, which is explained by the high viscosity and low surface tension of EPS and mucilage. During drying the polymers are progressively stretched and their viscosity exponentially increases until a critical point when the polymers can no longer follow the receding air-liquid interface. At this critical point the polymers start to behave as a solid matrix that enhances the soil water retention and connectivity of the liquid phase. A model is introduced to predict the formation these one- and two-dimensional polymer structures in drying porous media. Viscosity of the soil solution, drying rate and specific soil surface are the key parameters determining the transition from one- to two-dimensional structures. The model also explains the apparent hysteresis in the relation between water content and water potential in soils embedded with root mucilage. We propose that the release of viscous polymeric substances and the consequent creation of a network bridging the soil pore space represent a universal strategy of plants and bacteria to engineer their own soil micro-hydrological niches where stable conditions for life are preserved. [ABSTRACT FROM AUTHOR]

Plant roots and bacteria are capable of buffering erratic fluctuations of water content in their local soil environment by releasing a diverse, highly polymeric blend of substances (e.g. extracellular polymeric substances EPS and mucilage). Despite this concept is well accepted, the physical mechanisms by which EPS and mucilage interact with the soil matrix and determine the soil water dynamics remain unclear. High-resolution X-ray CT revealed that upon drying in porous media mucilage (from maize roots) and EPS (from intact biocrusts) form filaments and two-dimensional interconnected structures spanning across multiple pores. Unlike water, these mucilage and EPS structures connecting soil particles did not break up upon drying, which is explained by the high viscosity and low surface tension of EPS and mucilage. Measurements of water retention and evaporation with soils mixed with seed mucilage show how these one- and two-dimensional pore-scale structures impact macroscopic hydraulic properties: i.e. they enhance water retention, preserve the continuity of the liquid phase in drying soils and decreases vapor diffusivity and local drying rates. In conclusion, we propose that the release of viscous polymeric substances and the consequent creation of a network bridging the soil pore space represent a universal strategy of plants and bacteria to engineer their own soil micro-hydrological niches where stable conditions for life are preserved. [ABSTRACT FROM AUTHOR]

Background and Aims: Although maize roots have been extensively studied, there is limited information on the effect of root exudates on the hydraulic properties of maize rhizosphere. Recent experiments suggested that the mucilaginous fraction of root exudates may cause water repellency of the rhizosphere. Our objectives were: 1) to investigate whether maize rhizosphere turns hydrophobic after drying and subsequent rewetting; 2) to test whether maize mucilage is hydrophobic; and 3) to find a quantitative relation between rhizosphere rewetting, particle size, soil matric potential and mucilage concentration. Methods: Maize plants were grown in aluminum containers filled with a sandy soil. When the plants were 3-weeks-old, the soil was let dry and then it was irrigated. The soil water content during irrigation was imaged using neutron radiography. In a parallel experiment, ten maize plants were grown in sandy soil for 5 weeks. Mucilage was collected from young brace roots growing above the soil. Mucilage was placed on glass slides and let dry. The contact angle was measured with the sessile drop method for varying mucilage concentration. Additionally, capillary rise experiments were performed in soils of varying particle size mixed with maize mucilage. We then used a pore-network model in which mucilage was randomly distributed in a cubic lattice. The general idea was that rewetting of a pore is impeded when the concentration of mucilage on the pore surface (g cm) is higher than a given threshold value. The threshold value depended on soil matric potential, pore radius and contract angle. Then, we randomly distributed mucilage in the pore network and we calculated the percolation of water across a cubic lattice for varying soil particle size, mucilage concentration and matric potential. Results: Our results showed that: 1) the rhizosphere of maize stayed temporarily dry after irrigation; 2) mucilage became water repellent after drying. Mucilage contact angle increased with mucilage surface concentration (gram of dry mucilage per surface area); 3) Water could easily cross the rhizosphere when the mucilage concentration was below a given threshold. In contrast, above a critical mucilage concentration water could not flow through the rhizosphere. The critical mucilage concentration decreased with increasing particle size and decreasing matric potential. Conclusions: These results show the importance of mucilage exudation for the water fluxes across the root-soil interface. Our percolation model predicts at what mucilage concentration the rhizosphere turns hydrophobic depending on soil texture and matric potential. Further studies are needed to extend these results to varying soil conditions and to upscale them to the entire root system. [ABSTRACT FROM AUTHOR]

Here, we present our collective musings on soil research challenges and opportunities and, in particular, the points raised by Philippe Baveye (Baveye, 2020a, 2020b) and Johan Bouma (Bouma, 2020) on I bypass i and I hyperbole i in soil science. Furthermore, developing a healthy and constructive post-publication peer-review system, where bypasses and hyperbolic approaches can be identified and discussed, would ultimately boost publication quality and contribute to a more open discourse in soil science. [Extracted from the article]

Despite detailed investigations on the distinct biochemical properties of the rhizosphere and their effects on the availability of nutrients for plants, their biophysical aspects, particularly the effect of mucilage on the transport of water and nutrients are poorly understood. The aim of this study was to investigate the effect of mucilage on the diffusion coefficient of nutrients and consequently their transport in the soil and into the plant roots. Repeated phosphor imaging was used to monitor the temporospatial distribution of 137Cs (as an analog of K) within a modeled rhizosphere soil with and without mucilage (a sandy soil amended with mucilage extract from chia seed) under different soil water contents. The monitored profiles of activities were used to estimate the diffusion coefficient of soils with and without mucilage by solving a diffusion equation. Assuming an identical the effect of mucilage on diffusion coefficient of Cs and K, a diffusion-convection equation was numerically solved to predict the transport of K within the soil and its uptake by a single plant root during a soil drying cycle. To this end, the hydraulic and diffusive properties of the soil were parameterized based on the measured data and the K uptake and its concentration in soil were taken from literature data. The results of this study suggest that mucilage in the rhizosphere keeps the rhizosphere wetter and maintains the connectivity of the liquid phase during a soil drying cycle, and thereby could prevent a marked drop in the diffusion coefficient. The results of modeling of nutrient uptake by a single root showed that the presence of mucilage in the rhizosphere could favor nutrient uptake by the plant root. In the case of nutrients with low concentration in the soil solution, it prevents a marked concentration drop in the vicinity of the root as the soil dries and diffusion becomes restricted. This will delay the risk of nutrient deficiency to the plant root. In the case of nutrients with a high concentration in the soil solution, the presence of mucilage in the rhizosphere may delay the risk of salinity stress as the soil dries and the concentration of nutrients increases at the vicinity of the root surface. In conclusion, the results of this contribution show that mucilage may favor the transport of nutrient within the soil and their uptake by plant roots under drying soil condition. [ABSTRACT FROM AUTHOR]

Background and Aims: Recent experiments showed that rhizosphere of several plant species turns temporarily hydrophobic after severe drying and subsequent rewetting. Whether or not such hydrophobicity limits root water uptake is not known.Methods: A set of experiments was performed to test whether rhizosphere water repellency negatively affects root water uptake. To this end, a commercial surfactant was used as a rewetting agent to facilitate the wettability of the rhizosphere of lupins (Lupinus albus) in a sandy soil. Lupin plants were grown in rhizoboxes and were subjected to a severe drying cycle. Then half of the plants were irrigated with water and half with the surfactant solution. Time-series neutron radiography technique was used to monitor water redistribution in the rhizosphere during irrigation. In a second experimental set-up, lupins were grown in a sandy soil partitioned in five vertical compartments separated by a 1-cm layer of coarse sand (acting as a capillary barrier). Water and surfactant were injected in different compartments and the rehydration of the root tissues beyond the irrigated compartments was monitored with neutron radiography for 2-3 h. Root rehydration rates were used to estimate the water fluxes across the root-soil interface.Results: The rhizosphere of lupin roots in sandy soil irrigated with water remained partly dry for at least 2-3 h, while it was rapidly rewetted when irrigated with surfactant. Water flow into the roots irrigated with surfactant solution was 6.5 times faster than into the roots irrigated with water.Conclusions: These results prove that water repellency of the rhizosphere of lupins in sandy soils limited the water fluxes into the roots and root rehydration during the first two to three hours after irrigation. This might not always be negative, because it can limit water losses from roots to dry soil and therefore avoid severe root dehydration. [ABSTRACT FROM AUTHOR]